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The abundance and size distribution of plankton in the surface ocean are key metrics to understanding primary and export production, food web dynamics, and the optical properties of the water column. Here we quantified cell size and abundance of phytoplankton species from mono-specific laboratory cultures, using optical, electrical, and image-based benchtop instrumentation. Moreover, we combined particle size distribution and size fractionated chlorophyll a (Chl a) analysis with image analysis to estimate phytoplankton community composition and abundance comparing a high and low biomass station in the North Atlantic during springtime. In laboratory cultures, we found generally good agreement in estimates of both particle concentration and particle size among instruments. Image-based approaches (e.g., microscopy, FlowCam) delivered somewhat lower cell abundance estimates, because image-based instruments distinguish cells from non-target particles. Image-based approaches also measured 10–20% greater cell sizes, because measurements were based only on in-focus images of the target species. Compared to image-based instruments, particle counters delivered indistinguishable estimates of size and abundance with much lower effort and technical expertise required; maximum coefficient of variation for cell abundance and size did not exceed 10 and 15%, respectively. Measurement precision was consistent across instrument type and across a cell size spectrum from 3 to >40 μm equivalent spherical diameter (ESD). For whole plankton community analyses from the North Atlantic, combined estimation of Chl a concentration and image-based diversity assessments resolved the dominant phytoplankton species and spatial differences in the size structure of the plankton community. These results provide strong support for using particle counters, that can rapidly process large volumes of samples, to quantify particle size and abundance, including rarer, larger particles. Species identities and community composition can be revealed by supplemental, image-based approaches. Application of this coupled approach can help identify fundamental ecosystem characteristics such as particle size spectra that affect primary production, trophic transfer, and export. Ultimately, the tremendous species diversity of plankton can be leveraged as particle tracking and identification keys, such as near-real time identification of coherent water masses.

Phytoplankton growth dependence on temperature is recognized and has been quantified comprehensively. However, no similar relationship exists for the major phytoplankton predators, the herbivorous protists, especially at low temperatures representing polar and coastal oceans during most seasons. Their acclimation to changing temperatures is also largely unexplored. Here we report acclimated growth and acclimation rates from 0 to 22°C for 3 cosmopolitan herbivorous dinoflagellates. Due to interactive effects between size and temperature, growth increased 40% more rapidly with increasing temperature for production- compared to division-based growth rates (0.043 and 0.062 d-1 °C-1, respectively). Biomass-based growth rates were 10-fold higher than abundance-based rates at low temperatures, reflecting an average 50% increase in biovolume at ≤2°C. Thus, there was significant biomass accumulation at low temperatures, despite low cell-division rates. Testing different acclimation procedures, we established that acclimated rates emerged after 3 generations. Herbivores required 1.25 d °C-1 when acclimating towards higher temperatures and 2.5 d °C-1 when transitioning towards lower temperatures. Growth rates increased linearly with temperature, implying a weaker temperature effect on growth than the commonly assumed exponential dependency. A possible consequence is that herbivore growth rates are underestimated at cold and overestimated at warm temperatures. Current and future ocean assessments could thus underestimate trophic transfer rates in polar and cold-water regions and overestimate herbivore growth and thus grazing impact in future ocean predictions. Identifying physiological responses that transcend species-specificity supports cross-biome comparisons of ecosystem structure and function that rely on accurate predictions of matter and energy flow in planktonic food webs.

Diatoms produce a series of cytotoxic secondary metabolites such as polyunsaturated aldehydes (PUA) in response to cell injury or stress. However, little information exists on the PUA effects on the pelagic food web. A set of experiments was conducted in the Chesapeake Bay and the coastal Atlantic waters using dissolved PUA (2E,4E-octadienal and 2E,4E-heptadienal), natural assemblages of phytoplankton and microzooplankton, and the copepod Acartia tonsa. The results demonstrate that PUA primarily acts as deterrent for microzooplankton herbivory on diatoms, while enhancing herbivory on picophytoplankton. This switch should favor PUA-producing diatoms by simultaneously reducing direct grazing losses and competition. Additionally, PUA stimulated copepod predation on microzooplankton, particularly on ciliates. Combined, these effects have the potential to disrupt growth-grazing equilibrium in the pelagic food web and create an opportunity for bloom development.

During NAAMES May 2016 field campaign, we examined the response of plankton population dynamics to rapid re-stratification of surface waters following a deep mixing event in the subpolar North Atlantic. Over the 4-day occupation of a station, we measured phytoplankton growth and grazer/virus-induced mortality rates in dilution assays conducted across a range of light intensities. Rates were estimated from changes in chlorophyll a and the abundance of three phytoplankton groups identified by flow cytometry. Initially, biological and physical water column properties were homogeneous down to >200 m, followed by rapid shoaling of the mixed layer to 20-30 m. Initial in situ chlorophyll a concentration was 0.4 µg L-1, and phytoplankton biomass accumulated at an average rate of 0.4 d-1 over the next two days. When mixed layer depth was maximal, there were no mortality losses and phytoplankton growth rates increased with increased light intensity. After shoaling, grazing rates increased, but never matched the magnitude of phytoplankton growth rates. When the mixed layer was shallowest, growth rates exceeded >1 d-1 at all non-dark light intensities. Chlorophyll a based grazing rates were consistent across light levels (~0.3 d-1) and were highest on Synechococcus (0.3-0.6 d-1) and lowest on pico-eukaryotes (~0.2 d-1). The delay with which grazing resumed resulted in growth exceeding losses and consumption of an average of 30% of primary production. Virus-induced mortality rates were minimal across all mixing profiles and light intensities. Overall, both chlorophyll a and group-specific phytoplankton accumulation rates predicted from the shipboard experiments matched those in situ, suggesting that incubation conditions faithfully captured the growth and loss processes governing in situ population dynamics. The observations made here indicate that transient deepening of the mixed layer followed by rapid stratification provided conditions under which phytoplankton escape protistan grazing and accumulate biomass. During the physically dynamic springtime in the North Atlantic, frequent ephemeral fluctuations in the depth of mixing may represent an important mechanism governing the formation and the magnitude of the North Atlantic spring bloom.

Microzooplankton community structure, distribution, growth and herbivory were examined in the eastern Fram Strait and Arctic Ocean shelf affected by the Atlantic water inflow in May (during the spring bloom, chlorophyll up to 9 µg l-1) and August (post-bloom, 0.13 – 1.7 µg l-1) 2014. Microzooplankton were collected along two longitudinal transects at 79°N and 79.4 °N from the slope to the ice edge and crossed the Western Spitsbergen Current (WSC). In May, integrated microzooplankton biomass in the upper 100 m ranged from 0.16 g C m-2 above the slope to 2.3 g C m-2 within WSC (0.71 g C m-2 on average). Mixotrophic oligotrich ciliates from the genus Strombidium dominated in the spring and formed a surface bloom (79 x 103 cells L-1, 206 µg C L-1). This is the largest microzooplankton biomass recorded in the Arctic so far. The heterotrophic dinoflagellates Gyrodinium and Protoperidinium were abundant at the diatom-dominated stations in the marginal ice zone. In the summer, a more diverse community included a large proportion of heterotrophic and mixotrophic dinoflagellates, tintinnids, and other ciliates. Microzooplankton biomass increased to the average of 1.27 g C m-2. At the ice-covered and open water stations in the Yermak shelf and deep basin, microzooplankton grew at 0.04 to 0.38 d-1; their species-specific growth rates were up to 1.79 d-1. Microzooplankton herbivory on average removed 72% (36 to >100%) of daily primary production with the exception of Phaeocystis pouchetii colonies. These results indicate that the microbial food web plays a central role in the carbon cycle in this Atlantic-influenced polar system.

We used inverse modeling to reconstruct major planktonic food web carbon flows in the Atlantic Water inflow, east and north of Svalbard during spring (18-25 May) and summer (9-13 August), 2014. The model was based on three intensively sampled stations during both periods, corresponding to early, peak, and decline phases of a Phaeocystis and diatom dominated bloom (May), and flagellates dominated post bloom stages (August). The food web carbon flows were driven by primary production (290-2,850 mg C m d-1), which was channeled through a network of planktonic compartments, and ultimately respired (180-1200 mg C m d), settled out of the euphotic zone as organic particles (145-530 mg C m d), or accumulated in the water column in various organic pools. The accumulation of dissolved organic carbon was intense (1070 mg C m d) during the early bloom stage, slowed down during the bloom peak (400 mg C m d), and remained low during the rest of the season. The heterotrophic bacteria responded swiftly to the massive release of new DOC by high but decreasing carbon assimilation rates (from 534 to 330 mg C m d) in May. The net bacterial production was low during the early and peak bloom (26-31 mg C m d) but increased in the late and post bloom phases ( > 50 mg C m d). The heterotrophic nanoflagellates did not respond predictably to the different bloom phases, with relatively modest carbon uptake, 30-170 mg C m d. In contrast, microzooplankton increased food intake from 160 to 380 mg C m d during the buildup and decline phases, and highly variable carbon intake 46-624 mg C m d, during post bloom phases. Mesozooplankton had an initially high but decreasing carbon uptake in May (220-48 mg C m d), followed by highly variable carbon consumption during the post bloom stages (40-190 mg C m d). Both, micro- and mesozooplankton shifted from almost pure herbivory (92-97% of total food intake) during the early bloom phase to an herbivorous, detritovorous and carnivorous mixed diet as the season progressed. Our results indicate a temporal decoupling between the microbial and zooplankton dominated heterotrophic carbon flows during the course of the bloom in a highly productive Atlantic gateway to the Arctic Ocean., This work was funded by the Estonian Research Council (Grant 1574P), and the Norwegian Research Council through the project CarbonBridge (Project Number 226415).

Source at https://doi.org/10.3389/fmars.2019.00293. The Arctic Ocean is changing rapidly with respect to ice cover extent and volume, growth season duration and biological production. Zooplankton are important components in the arctic marine food web, and tightly coupled to the strong seasonality in primary production. In this study, we investigate zooplankton composition, including microzooplankton, copepod nauplii, as well as small and large copepod taxa, and primary productivity in the dynamic Atlantic water inflow area north of Svalbard in May and August 2014. We focus on seasonal differences in the zooplankton community and in primary productivity regimes. More specifically, we examine how a shift from “new” (nitrate based) spring bloom to a “regenerated” (ammonium based) post bloom primary production is reflected in the diversity, life history adaptations and productivity of the dominant zooplankton. North of Svalbard, the seasonal differences in planktonic communities were significant. In spring, the large copepod Calanus finmarchicus dominated, but the estimated production and ingestion rates were low compared to the total primary production. In summer, the zooplankton community was composed of microzooplankton and the small copepod Oithona similis. The zooplankton production and ingestion rates were high in summer, and probably depended heavily on the regenerated primary production associated with the microbial loop. There was clear alteration from dominance of calanoid copepod nauplii in spring to Oithona spp. nauplii in summer, which indicates different reproductive strategies of the dominating large and small copepod species. Our study confirms the dependence and tight coupling between the new (spring bloom) primary production and reproductive adaptations of C. glacialis and C. hyperboreus. In contrast, C. finmarchicus appears able to take advantage of the regenerated summer primary production, which allows it to reach the overwintering stage within one growth season in this region north of Svalbard. This suggests that C. finmarchicus will be able to profit from the predicted increased primary production in the Arctic, a strategy also recognized in small copepod species such as O. similis. We speculate that the ability of the copepod species to utilize the regenerated summer primary production and microbial food web may determine the winners and losers in the future Arctic Ocean.

Allelopathy is wide spread among marine phytoplankton, including diatoms, which can produce cytotoxic secondary metabolites such as polyunsaturated aldehydes (PUA). Most studies on diatom-produced PUA have been dedicated to their inhibitory effects on reproduction and development of marine invertebrates. However, little information exists on their impact on key herbivores in the ocean, microzooplankton. This study examined the effects of dissolved 2E,4E-octadienal and 2E,4E-heptadienal on the growth rates of natural ciliate and dinoflagellate populations in the Chesapeake Bay and the coastal Atlantic waters. The overall effect of PUA on microzooplankton growth was negative, especially at the higher concentrations, but there were pronounced differences in response among common planktonic species. For example, the growth of Codonella sp., Leegaardiella sol, Prorodon sp., and Gyrodinium spirale was impaired at 2 nM, whereas Strombidium conicum, Cyclotrichium gigas, and Gymnodinium sp. were not affected even at 20 nM. These results indicate that PUA can induce changes in microzooplankton dynamics and species composition.

The internal organization of plankton communities plays a key role in biogeochemical cycles and in the functioning of aquatic ecosystems. In this study, the structure of a marine plankton community (including both unicellular and multicellular organisms) was inferred by applying an ecological network approach to species abundances observed weekly at the long-term ecological research station MareChiara (LTER-MC) in the Gulf of Naples (Tyrrhenian Sea, Mediterranean Sea) in the summers of 2002–2009. Two distinct conditions, characterized by different combination of salinity and chlorophyll values, alternated at the site: one influenced by coastal waters, herein named ‘green’, and the other reflecting more offshore conditions, named ‘blue’. The green and blue ‘phases’ showed different keystone biological elements: namely, large diatoms and small-sized flagellates, respectively. Several correlations amongst species belonging to different trophic groups were found in both phases (connectance ~0.30). In the green phase, several links between phytoplankton and mesozooplankton and within the latter were detected, suggesting matter flow from microbes up to carnivorous zooplankton. A microbial-loop-like sub-web, including mixo- and heterotrophic dinoflagellates and ciliates, was present in the green phase, but it was relatively more important in the blue phase. The latter observation suggests a more intense cycling of matter at the microbial trophic level in the blue phase. These results show that different modes of ecological organization can emerge from relatively small changes in the composition of aquatic communities coping with environmental variability. This highlights a significant plasticity in the internal structure of plankton webs, which should be taken into account in predictions of the potential effects of climatic oscillations on aquatic ecosystems and biogeochemical cycles therein.

Invasive quagga mussels have recently replaced zebra mussels as the dominant filter-feeding bivalves in the Great Lakes. This study examined microzooplankton (i.e., grazers Microcystis and diatom blooms, respectively. Feeding on microzooplankton could buffer mussels during lean periods, or supplement other consumed resources, particularly during noxious cyanobacterial blooms. The results of this study demonstrate that microzooplankton are a resilient and critical component of the Saginaw Bay ecosystem.

The fate of microzooplankton production, whether it is channeled to mesozooplankton or recycled within the microbial food web, has major implications for the oceanic carbon cycle. The aim of this study was to estimate internal predation within naturally occurring microzooplankton communities. A dilution series based on the Landry and Hasset technique was created by mixing 200-μm-screened water (used as whole water) with 5-μm-screened seawater due to the dominance of pico- and small nanoplankton at our study site. This modification of the original technique allows for gradual reduction in microzooplankton abundance and thus internal predation while maintaining sufficient phytoplankton prey levels for microzooplankton growth in diluted treatments. Microzooplankton growth and mortality rates were calculated based on the changes in abundance during 24-h incubation. In the diluted treatments, microzooplankton growth rates were significantly higher (1.21 ± 0.20 day−1 for ciliates and 0.88 ± 0.05 day−1 for heterotrophic dinoflagellates) compared to those in whole seawater where microzooplankton abundance remained unchanged or even declined over time. Approximately 79 % of microzooplankton production was consumed within the microzooplankton, with aloricate ciliates being the most vulnerable to predation. These findings support the assumption that trophic interactions between microzooplankton can be an important factor controlling their production and, thus, energy transfer in picoplankton-dominated pelagic ecosystems.

Microzooplankton grazing on phytoplankton populations was investigated by means of eight dilution experiments in different seasons and trophic conditions at station Mare Chiara, a longterm study site in the Gulf of Naples, Tyrrhenian Sea. To check for changes in prey and predator populations, size-fractionated chlorophyll and HPLC analyses and microzooplankton counts were performed before and after the incubations. On average, 68% of daily phytoplankton production was consumed by the microzooplankton in spring and summer, and in winter, more than 100% of phytoplankton daily production was consumed. Nutrients were added in four of our experiments. Diatoms (as recorded by fucoxanthin concentrations) increased significantly in the nutrient-enriched experiments, and unexpectedly, also heterotrophic dinoflagellate growth was enhanced by nutrient additions. Remarkable changes in grazer abundance were observed in several experiments; a substantial difference was found in microzooplankton grazing pressure (g/k � 100) as calculated by regression of apparent chlorophyll a growth against nominal dilution factor or against average predator concentrations, 72+19% and 60+23% of phytoplankton daily production, respectively. However, our observations suggest that if growth and mortality of grazers in the dilution series are not considered, microzooplankton grazing impact may be either underestimated, at low phytoplankton growth rates, or overestimated in more productive conditions.

Growth rates (µ) of abundant microzooplankton species were examined in field experiments conducted at ambient sea temperatures (-1.8-9.0°C) in the Barents Sea and adjacent waters (70-78.5°N). The maximum species-specific µ of ciliates and athecate dinoflagellates (0.33-1.67 d(-1) and 0.52-1.14 d(-1), respectively) occurred at temperatures below 5°C and exceeded the µmax predicted by previously published, laboratory culture-derived equations. The opposite trend was found for thecate dinoflagellates, which grew faster in the warmer Atlantic Ocean water. Mixotrophic ciliates and dinoflagellates grew faster than their heterotrophic counterparts. At sub-zero temperatures, microzooplankton µmax matched those predicted for phytoplankton by temperature-dependent growth equations. These results indicate that microzooplankton protists may be as adapted to extreme Arctic conditions as their algal prey.